Tropics 16-1.Indb

TROPICS Vol. 16 (1) Issued January 31, 2007

Fruit visitation patterns of small mammals on the forest floor in a tropical seasonal forest of Thailand

1 2, 3 1 3 3 3 Shunsuke SUZUKI , Shumpei KITAMURA , Masahiro KON , Pilai POONSWAD , Phitaya CHUAILUA , Kamol PLONGMAI , Takakazu 2, 4 1 5 6 YUMOTO , Naohiko NOMA , Tamaki MARUHASHI and Prawat WOHANDEE

1 School of Environmental Science, The University of Shiga Prefecture, Hikone, 522−8533, Japan 2 Center for Ecological Research, Kyoto University, Kamitanakami-Hirano, Otsu, 520−2113, Japan 3 Thailand Hornbill Project, Department of Microbiology, Faculty of Science, Mahidol University, Bangkok 10400, Thailand 4 Research Institute of Humanity and Nature, 457-7, Motoyama, Kamigamo, Kita-ku, Kyoto, 602−0878, Japan 5 Department of Human and Culture, Musashi University, Nerima, Tokyo 176−8534, Japan 6 National Park, Wildlife and Plant Conservation Department, Bangkok 10900, Thailand

Correspondence address: Shunsuke SUZUKI Tel: +81−749−28−8311, Fax: +81−749−28−8477, E-mail: [email protected]

ABSTRACT The fruit visitation patterns of small birds, (Harrison, 1962; Miura et al. 1997), including mammals were investigated by camera trappings small mammals (Robinson et al. 1995; Wu et al. 1996; on the forest floor in a tropical seasonal forest of Yasuda et al. 2003; Pardini, 2004). The species richness Thailand. A total of 3,165 visits were recorded for and abundance of small mammals in tropical forests can seven small mammal species. The four Muridae be attributed to affluent food resources, especially the species, Rattus remotus, Niviventer fulvescens, abundant fruit crops (Fleming, 1973; August, 1983; Wells Leopoldamys sabanus and Maxomys surifer , all et al. 2004). Terrestrial small mammals are considered of which were nocturnal, were almost completely to have effects on tropical forest ecosystems as seed temporally segregated from the tree shrew, Tupaia and seedling predators and seed dispersers (Adler and belangeri , and the two squirrels, Callosciurus Kestell, 1998; Asquith et al. 1999; Guariguata et al. 2000). finlaysonii and Menetes berdmorei , which were In recent years, the abundance of large animal species diurnal or crepuscular. We suggest that the has declined and communities have been altered due to temporal segregation reduced the interference habitat loss or fragmentation in tropical forests (Soule et competition between the four Muridae species and al. 1992; Laidlaw, 2000; Sodhi et al. 2004). Small mammal the tree shrew or squirrel for fruits on the forest species, however, are adaptable enough to inhabit in floor. In addition, the visitation patterns for fruit various and patchy environments and are more robust species differed among the four Muridae species in dealing with habitat changes. In order to predict the and between the tree shrew and the two squirrels, fate of fragmented or disturbed forests, it is important to suggesting that the variation in the visitation understand the coexistence patterns of small mammal patterns for fruit species helped to facilitate assemblages in relation to their roles in tropical forest coexistence among these species. In contrast, ecosystems (Wells et al. 2004). the two squirrels were similar in their visitation The coexistence of sympatric species can be patterns, both temporally and in their choice of facilitated by the differentiation in niche dimensions, fruit species. such as habitat, food and time (Schoener, 1974; , M Closkey, 1976). There have been several studies on Key words: coexistence, crepuscular, diurnal, microhabitat utilization and segregation of small mammal frugivore, Khao Yai National Park, nocturnal, small assemblages in tropical forests (Shanker, 2001; Wells mammal, temporal segregation et al. 2004). While there have been some studies on food resource partitioning in rodents in the desert (e.g. Brown et al. 1979; Brown, 1989), there have been very INTRODUCTION few on small mammals in tropical forests (Emmons, 1980; Tropical forests are enormously diverse and complex Smythe, 1986). Moreover, little is known about temporal habitats and harbor many species of mammals and differentiation of food resource utilization. This is 18 Shunsuke SUZUKI, Shumpei KITAMURA, Masahiro KON, Pilai POONSWAD, Phitaya CHUAILUA, Kamol PLONGMAI, Takakazu YUMOTO, Naohiko NOMA, Tamaki MARUHASHI and Prawat WOHANDEE

partly due to the difficulty in observing the often cryptic Park in Thailand. We obtained a large number of pictures behavior of small mammals. Temporal differentiation of small mammals visiting the fruit bait on the forest in food resource utilization can be recorded by various floor. In this paper, we examine the differentiation in the methods, including radio telemetry, consecutive fruit visitation patterns among small mammal species trapping, direct observation and camera trapping. In that utilize fruits on the forest floor, and compare the recent years, remarkable improvements have been made distribution of these patterns between small mammal for various wildlife studies through the use of camera species, both temporally and in terms of fruit species. trapping (Griffiths and van Schaik, 1993; Miura et al. 1997; Blanchong and Smale, 2000; Jayasekara et al. , 2003; O Brien et al. 2003; Kawanishi and Sunquist, 2004; MATERIALS AND METHODS Yasuda et al. 2005). Camera trapping can provide precise Study site time data for visitation patterns of animals, which can be The study was carried out in Khao Yai National Park quantitatively analyzed for visitation time and resource (hereafter KY; Fig. 1) in lower northeastern Thailand. , , utilization patterns (Yasuda et al. 2005). The park ranges from 14˚05 N to 14˚15 N and from 101˚ , , Through the use of camera trappings, the 05 E to 101˚ 50 E in the Dongruk mountain range, and 2 relationships between fruits and frugivores on the forest covers an area of 2,168 km . Its elevation ranges from floor of a tropical forest were studied in Khao Yai National 250 to 1,351 m. The main study site was adjacent to the 2 headquarters of KY and covered about 70 km . The area ranges from 600 to 800 m in altitude and is covered by moist evergreen forest. The annual mean rainfall is 2,360 mm (1993−2002); the wet season usually occurs from April to October and the dry season is from November to March (Kitamura et al. 2004). The monthly mean temperature ranges from 21˚C (December and January) to 32˚C (April and May). In the study area, sunrise is between 0545 h and 0643 h throughout a year and sunset is between 1743 h and 1847 h. According to the standard time near KY, we �� divided a day into the following three time zones: daytime �� from 0700 h to 1700 h; nighttime from 1900 h to 0500 h; and crepuscular time from 0500 h to 0700 h and 1700 h to 1900 h. �� Although ripe fruits (e.g. Ficus spp.) are available year-round (Poonswad et al. 1998), fruit diversity and abundance are relatively high in the rainy season and reach a trough at the beginning of the dry season (Bartlett 2003). Trees reach 45 m in height; the density of trees over 10 cm in diameter at breast height (DBH) is 371 ha−1, with a basal area of 32 m2ha−1. The dominant plant families in the forest include the Lauraceae, Cornaceae, Euphorbiaceae, Meliaceae, Dipterocarpaceae, and Annonaceae. Kitamura et al. (2005) provided a detailed botanical inventory.

Photographing by automatic camera systems We conducted surveys on the fruit visitation patterns of small mammals from July 2000 to June 2002 using automatic camera systems. We used two kinds of systems, both of which consist of a far-infrared sensor, Fig. 1. Location of Khao Yai National Park motor-driven compact camera with a built-in flash and Fruit visitation patterns of small mammals in Thailand 19 a data pack that stamps each picture with the date and based only on the data for the fruit species that had been time of exposure. The system used until June 2001 was visited by both animal species. developed by Miura et al. (1997). From April 2001, we used a Sensor Camera“ FIELDNOTE” (MARIF Co. Ltd, Similarity in the visitation pattern for fruit species Japan; Yasuda, 2004). The cameras were checked at an Similarity in the rank order for the number of visits per interval of 1 to 3 days, and ﬁlms (ISO 100 or ISO 400, 36 day to a fruit species was measured for each pair of , exposures) and batteries (CR123A lithium battery) were animal species using Spearman s rank correlation. If renewed if necessary. both of two given animal species have the same rank We used the fruits of 186 individuals of 67 plant order, then the rank correlation coefﬁcient culminates in species as baits. Five to 50 fruits that had been collected 1; if animals have a completely inverse rank order, it is −1; from the ground or trees were set on the ground under and if independent of each other, it is 0. the source tree as bait, and a camera was placed about 2 m apart from the bait fruits. When the bait fruits were consumed by animals or damaged by insects or microbes, RESULTS they were replaced with new ones. The photographing Forty-eight animal species were identified from 11,079 duration varied from 5 to 90 consecutive days according pictures. Among them, eight small mammal species to the period when fruits were available from each were included, but one species (Crocidura horsfildi) was individual tree. Based on the pictures, we recorded excluded from further analyses due to the small sample when and what small mammals visited the bait fruits. size (< 10 visits). The remaining seven species were We followed Corbet and Hill (1992) for the taxonomic recognized from 5,419 pictures. From the pictures of nomenclature of small mammals. small mammal species, 3,165 (58.4%) were regarded as Because the cameras were triggered every 10 to 15 independent visits and used for further analyses. Of seconds while an animal or group of animals was within these species, the yellow rajah rat, Maxomys surifer, the detectable area of the sensor, one picture does not visited most frequently, followed by the Indochinese necessarily represent a single visit of a given animal ground squirrel, Menetes berdmorei (Table 1). The visits species. We regarded a picture as one visit by a given of these two species accounted for 82.4 % of all the visits species only when the picture was taken more than 30 by the target species. , , minutes after the last picture of the same species (O Brien The Finlayson s squirrel, Callosciurus finlaysonii, et al. 2003, Yasuda, 2004). visited the bait fruits mainly in the daytime with a peak in the afternoon; about 90 % of its visits were recorded in Similarity in the temporal distribution of visits the daytime whereas there were no visits in the nighttime

Similarity in the temporal distribution of visits between a (Fig. 2a, Table 1). Hereafter we regard this species as , , , given pair of animal species was measured using Pianka s diurnal . In contrast, the Indochinese ground squirrel, α index as follows (Pianka, 1973): Menetes berdmorei, and the northern tree shrew, Tupaia belangeri, visited bait fruits bimodally with peaks (more Σpij pik i αjk= 2 2 than 50 % of their visits) at dawn and dusk crepuscular √ Σ‾‾‾‾‾‾(pij ) Σ(pik )

i i time (Fig. 2b and c, Table 1). Hereafter we regard these , , where pij is the proportion of visits during an hour i (i=0 species as crepuscular . All four Muridae species visited

− 23 h) to the total visits for animal species j and pik is that the bait fruits mainly in the nighttime when more than for animal species k. This index ranges between 0 and 1, 90 % of their visits were recorded, and there were no

and similarity increases as the index approaches 1. visits in the daytime (Fig. 2d − g, Table 1). Hereafter we , , Not all small mammal species investigated in the regard these species as nocturnal . present study necessarily utilized all the fruit species. Similarities in the temporal distribution of visits were However, if both of two given mammal species were low (α = 0.000 − 0.043) for species pairs between the four observed to visit a given fruit species, we considered that Muridae species and the tree shrew or squirrels (Table they potentially competed with each other for the fruit 2), whereas they were high for a pair of crepuscular species (although the intensity of competition could have species and those of nocturnal species (α = 0.678 − 0.983). changed with the relative abundance of the fruit species). Intermediate values were observed between diurnal Therefore, similarity in the temporal distributions of and crepuscular species (α = 0.398 for Tupaia belangeri visits between a given pair of animal species was analyzed and Callosciurus finlaysonii; α = 0.478 for Callosciurus 20 Shunsuke SUZUKI, Shumpei KITAMURA, Masahiro KON, Pilai POONSWAD, Phitaya CHUAILUA, Kamol PLONGMAI, Takakazu YUMOTO, Naohiko NOMA, Tamaki MARUHASHI and Prawat WOHANDEE

Table 1. Classification of small mammals based on the temporal distribution of visits. No. of Total Crepuscula Family Scientific name Daytime Nighttime Category pictures visits r time Sciuridae Callosciurus finlaysonii 307 106 99 (93.4) 7 (6.6) 0 (0.0) Diurnal Menetes berdmorei 1119 527 209 (39.7) 318 (60.3) 0 (0.0) Crepuscular

Tupaiidae Tupaia belangeri 192 121 54 (44.6) 67 (55.4) 0 (0.0) Crepuscular Muridae Rattus remotus 85 63 0 (0.0) 3 (4.8) 60 (95.2) Nocturnal Niviventer fulvescens 278 194 0 (0.0) 17 (8.8) 177 (91.2) Nocturnal Leopoldamys sabanus 120 73 0 (0.0) 0 (0.0) 73 (100.0) Nocturnal Maxomys surifer 3318 2081 0 (0.0) 134 (6.4) 1947 (93.6) Nocturnal total of small mammals 5419 3165 362 (0.0) 546 (6.4) 2257(93.6) No. of pictures: the total number of pictures of each small mammal species Total visits: the total number of visits Daytime: the number of visits recorded during the daytime Crepuscular time: the number of visits recorded during the crepuscular time Nighttime: the number of visits recorded during the nighttime The percentage to the total number of visits is shown in the parenthesis.

finlaysonii and Menetes berdmorei; Fig. 3a). their visitation patterns for each fruit species. Therefore, Fig. 3 shows the number of visits per day of each the differentiation in the preference of fruit species small mammal species for the representative fruit species. could have promoted coexistence among these species. The fruit species most frequently visited differed among However, we note that the fruit species visitation pattern small mammal species. Moreover, the rank orders for the recorded in the present study may not represent the number of visits to a fruit species per day were not highly resource utilization curve of a given mammal species, positively correlated for most pairs of animals (rs = −0.031 because we had no data on the relative abundance of each to 0.359; Table 2). The highest correlation was observed fruit species. between Tupaia belangeri and Rattus remotus (rs = 0.359, The two squirrel species more or less resembled P<0.01, N=63), followed by T. belangeri and Leopoldamys each other in both their temporal distribution of visits sabanus (rs = 0.268, P<0.05, N=63) and Callosciurus and visitation patterns for each fruit species (Table 2). finlaysonii and Menetes berdmorei (rs = 0.251, P<0.05, However, C. finlaysonii usually inhabits the middle storey N=63). There was no significant correlation between any of a forest canopy and is rarely found on the ground the other pairs of animal species. (Srikosamatara and Hansel, 1996), whereas M. berdmorei stays mainly on the ground (Lekagul and McNeely, 1977). Thus, these two squirrel species may be segregated DISCUSSION vertically rather than temporally or for fruit species. The temporal distribution of visits observed in the The ground squirrel, Menetes berdmorei, and the tree present study are regarded as representing the food shrew, Tupaia belangeri, were crepuscular in this study searching activity pattern of each small mammal species, and are known to forage mainly on the ground (Lekagul because we recorded their visits by attracting them with and McNeely, 1977; Emmons, 2000). However, they bait fruits. Differentiation in the temporal distribution appeared to be segregated in their visitation pattern for of visits between the four Muridae species and the other fruit species (Table 2, Figs. 2b and c). Menetes berdmorei three diurnal or crepuscular species is likely to have frequently visited hard covered fruits such as Canarium contributed to a reduction in interference competition euphyllum, Lithocarpus thomsonii, and Beilschmiedia for fruits on the forest floor. Moreover, if fruits supply villosa, whereas the tree shrew rarely visited them. It is limited, the temporal differentiation in food searching could be because M. berdmorei has well-developed incisor activities could also affect exploitative competition teeth whereas T. belangeri does not. In addition, T. between these species, depending on the timing of when belangeri also differed from another squirrel species, C. fruits fall to the ground. finlaysonii, in its visitation pattern for fruit species (Table The four Muridae species resembled each other 2). This appeared partly because C. finlaysonii also closely in their temporal distribution of visits, but not in visited C. euphyllum frequently. Fruit visitation patterns of small mammals in Thailand 21

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Fig. 2. Temporal distribution of visits by the seven small mammal species. Open column: daytime, gray column: crepuscular time, closed column: nighttime. 22 Shunsuke SUZUKI, Shumpei KITAMURA, Masahiro KON, Pilai POONSWAD, Phitaya CHUAILUA, Kamol PLONGMAI, Takakazu YUMOTO, Naohiko NOMA, Tamaki MARUHASHI and Prawat WOHANDEE

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Fig. 3. The number of visits per day of each small mammal specie for representative fruit species, including the three fruit species that each mammal species visited most frequently. 1. Alphonsea cf. cylindrica, 2. Miliusa lineata, 3. Canarium euphyllum, 4. Mastixia pentandra, 5. Euphorbiaceae sp., 6. Lithocarpus thomsonii, 7. Gonocarium lobbianum, 8. Beilschmiedia villosa, 9. Cryptocarya cf. impressa, 10. Phoebe cathia, 11. Acrocarpus fraxinifolius, 12. Aglaia lawii, 13. Ficus benjamina, 14. Ficus subcordata, 15. Anthocephalus chinensis, 16. Canthium coffeoides, 17. Citrus limon, 18. Mischocarpus pentapetalus. Fruit visitation patterns of small mammals in Thailand 23 , Table 2. Similarities between small mammals in the temporal distribution of visits ,measured by Pianka s α (A) and those in visitation pattern for fruit species measured by Spearman s rank correlation (B).

(A) Similarities in temporal distribution of visits (see Appendix for the raw data) Cf Mb Tb Rr Nf Ls Ms (19) (33) (28) (14) (23) (19) (59) Cf 15 10 5 9 9 19 Mb 0.478 18 10 16 12 32 Tb 0.398 0.895 10 12 12 26 Rr 0.000 0.017 0.017 5 7 12 Nf 0.000 0.018 0.017 0.678 9 23 Ls 0.000 0.000 0.000 0.794 0.908 18 Ms 0.002 0.043 0.031 0.952 0.983 0.925

(B) Similarities in the visitation pattern for fruit species (see Table 3 for the raw data) Cf Mb Tb Rr Nf Ls Ms (19) (33) (28) (14) (23) (19) (59) Cf 15 10 5 9 9 19 ＊ Mb 0.251 18 10 16 12 32 Tb 0.062 0.146 10 12 12 26 ＊＊ Rr 0.068 0.216 0.359 5 7 12 Nf 0.097 0.234 0.095 −0.031 9 23 ＊ Ls 0.176 0.099 0.268 0.231 0.048 18 Ms 0.043 0.224 0.093 0.170 0.113 0.109 The total number of fruit species visited by each mammal species is shown in the parenthesis on the ﬁrst line. Above diagonal: the number of fruit species visited by both the mammal species of each pair; below diagonal: the similarity in the activity pattern (A) and that in the fruit species visit pattern (B). ＊＊＊ : p<0.01, p<0.05. Cf: Callosciurus finlaysonii, Mb: Menetes berdmorei, Tb: Tupaia belangeri, Rr: Rattus remotus, Nf: Niviventer fulvescens, Ls: Leopoldamys sabanus, Ms: Maxomys surifer.

The present results suggest that the differentiation Hornbill Project for supporting our fieldwork and in the visitation patterns for fruit species and feeding providing encouragement and hospitality. We extend time promoted coexistence between some pairs of our hearty thanks to the staffs of KY. This research small mammals in the tropical forest of KY. However, was supported in part by a Research Fund of the Japan it is known that most small mammal species (including Society for the Promotion of Science (#1357006) and rodents, squirrels and tree shrews) often feed on JSPS Research Fellowships for Young Scientists for S. arthropods as well as fruits (Harrison, 1954, 1961; Kitamura. Lekagul and McNeely, 1977; Corbet and Hill, 1992; Srikosamatara and Hansel, 1996; Emmons, 2000). Thus, further studies on arthropod utilization patterns REFERENCES by small mammals are necessary to understand more Adler, G.H. & Kestell D.W. 1998. Fates of neotropical comprehensively the factors facilitating coexistence tree seeds influenced by spiny rats (Proechimys among small mammals. semispinosus). Biotropica, 30: 677−681. Asquith, N.M., Terborgh J., Arnold A.E. & Riveros C.M. ACKNOWLEDGMENTS We are grateful to the 1999. The fruits the agouti ate: Hymenaea courbaril National Research Council of Thailand and the National seed fate when its disperser is absent. Journal of Park Division of the Royal Forest Department of Thailand Tropical Ecology, 15: 229−235. for allowing us to conduct our research in KY. We August, P.V. 1983. The role of habitat complexity and thank B. Saengthong, S. Chuailua, S. Nakkuntod, S. heterogeneity in structuring tropical mammal Sanguanchat, N. Jirawatkavi and all staff of the Thailand communities. Ecology, 64: 1495−1507. 24 Shunsuke SUZUKI, Shumpei KITAMURA, Masahiro KON, Pilai POONSWAD, Phitaya CHUAILUA, Kamol PLONGMAI, Takakazu YUMOTO, Naohiko NOMA, Tamaki MARUHASHI and Prawat WOHANDEE 6 ( 0 . 171 ) 1 ( 0 . 053 ) 2 ( 0 . 167 ) 6 ( 0 . 188 ) 5 ( 0 . 094 ) 3 ( 0 . 143 ) 4 ( 0 . 111 ) 1 ( 0 . 017 ) 9 ( 0 . 391 ) 7 ( 0 . 304 ) surifer 18 ( 0 . 667 ) 12 ( 0 . 088 ) 37 ( 1 . 000 ) 22 ( 0 . 917 ) 37 ( 0 . 544 ) 21 ( 0 . 233 ) 53 ( 0 . 510 ) 12 ( 0 . 185 ) 57 ( 1 . 390 ) 11 ( 0 . 478 ) 66 ( 0 . 528 ) 28 ( 0 . 718 ) 17 ( 1 . 000 ) 16 ( 0 . 471 ) 40 ( 0 . 667 ) 26 ( 0 . 245 ) 78 ( 1 . 444 ) 32 ( 0 . 252 ) Maxomys 155 ( 0 . 525 ) 227 ( 1 . 201 ) 159 ( 1 . 544 ) 1 ( 0 . 003 ) 8 ( 0 . 229 ) 5 ( 0 . 185 ) 5 ( 0 . 037 ) 1 ( 0 . 042 ) 4 ( 0 . 059 ) 1 ( 0 . 010 ) 7 ( 0 . 206 ) sabanus 24 ( 0 . 127 ) Leopoldamys 4 ( 0 . 014 ) 2 ( 0 . 054 ) 7 ( 0 . 037 ) 2 ( 0 . 019 ) 2 ( 0 . 038 ) 5 ( 0 . 048 ) 9 ( 0 . 250 ) 4 ( 0 . 103 ) 2 ( 0 . 059 ) 1 ( 0 . 017 ) 6 ( 0 . 047 ) 8 ( 0 . 348 ) fulvescens Niviventer Rattus 2 ( 0 . 007 ) 1 ( 0 . 029 ) 8 ( 0 . 296 ) 1 ( 0 . 005 ) 4 ( 0 . 059 ) 2 ( 0 . 051 ) remotus 10 ( 0 . 417 ) Tupaia 3 ( 0 . 086 ) 2 ( 0 . 074 ) 1 ( 0 . 007 ) 7 ( 0 . 037 ) 1 ( 0 . 010 ) 1 ( 0 . 011 ) 6 ( 0 . 058 ) 3 ( 0 . 214 ) 1 ( 0 . 059 ) 1 ( 0 . 009 ) 1 ( 0 . 019 ) belangeri 14 ( 0 . 047 ) 12 ( 0 . 500 ) 8 ( 0 . 229 ) 5 ( 0 . 135 ) 9 ( 0 . 087 ) 1 ( 0 . 019 ) 1 ( 0 . 015 ) 8 ( 0 . 064 ) 3 ( 0 . 125 ) Menetes 21 ( 0 . 071 ) 41 ( 0 . 394 ) 32 ( 1 . 391 ) 10 ( 0 . 256 ) 20 ( 0 . 333 ) 19 ( 0 . 826 ) 11 ( 0 . 104 ) 16 ( 0 . 126 ) berdmorei 254 ( 1 . 344 ) 5 ( 0 . 017 ) 1 ( 0 . 007 ) 1 ( 0 . 083 ) 1 ( 0 . 008 ) 1 ( 0 . 017 ) 6 ( 0 . 057 ) 2 ( 0 . 016 ) finlaysonii 30 ( 0 . 159 ) Callosciurus 35 27 19 37 24 12 32 68 53 21 90 65 36 14 41 23 58 39 17 34 60 23 54 24 23 295 136 189 103 104 125 106 127 Day sp. sp. sp. sp. Speceis Name Choerospondias axillaris Alphonsea elliptica Miliusa cuneata Platymitra macrocarpa Polyalthia jucunda Polyalthia Polyalthia viridis Uvaria lurida Canarium euphyllum Bhesa robusta Mastixia pentandra tricuspidata Trichosanthes Dipterocarpus gracilis Diospyros glandulosa Elaeagnus conferta Elaeocarpus robstus Balakata baccata Macaranga gigantea Unidentified(DCT# 233 ) Lithocarpus Lithocarpus thomsonii Quercus myrsinaefolia Casearia grewiaefolia Gnetum Garcinia merguensis Gonocarium lobbianum Platea latifolia Beilschmiedia villosa Cinnamomum subavenium Cryptocarya impressa Phoebe cathia Acacia Acrocarpus fraxinifolius Family Anacardiaceae Annonaceae Burseraceae Celastraceae Cornaceae Cucurbitaceae Dipterocarpaceae Ebenaceae Elaeagnaceae Elaeocarpaceae Euphorbiaceae Fagaceae Flacourtiaceae Gnetaceae Guttiferae Icacinaceae Lauraceae Leguminosae . The number of visits seven small mammal species to each fruit speceis. Table 3 . Fruit visitation patterns of small mammals in Thailand 25 ( 0 . 051 ) ( 0 . 051 ) 6 ( 0 . 500 ) 8 ( 0 . 088 ) 7 ( 0 . 179 ) 6 ( 0 . 222 ) 5 ( 0 . 417 ) 8 ( 0 . 444 ) 5 ( 0 . 096 ) 4 ( 0 . 154 ) 7 ( 0 . 500 ) 3 ( 0 . 051 ) surifer 11 ( 0 . 611 ) 22 ( 0 . 500 ) 19 ( 0 . 514 ) 22 ( 0 . 214 ) 33 ( 0 . 660 ) 46 ( 0 . 338 ) 65 ( 0 . 546 ) 55 ( 0 . 797 ) 34 ( 1 . 360 ) 10 ( 0 . 500 ) 33 ( 0 . 248 ) 40 ( 0 . 471 ) 35 ( 0 . 745 ) 89 ( 1 . 435 ) 21 ( 0 . 488 ) 60 ( 1 . 935 ) 18 ( 0 . 247 ) 60 Maxomys 241 ( 1 . 296 ) 2081 ( 0 . 051 ) ( 0 . 051 ) 1 ( 0 . 010 ) 1 ( 0 . 011 ) 3 ( 0 . 097 ) 2 ( 0 . 017 ) 1 ( 0 . 040 ) 2 ( 0 . 100 ) 3 ( 0 . 064 ) 1 ( 0 . 016 ) 1 ( 0 . 023 ) 2 ( 0 . 034 ) sabanus 20 73 Leopoldamys ( 0 . 051 ) ( 0 . 051 ) 3 ( 0 . 029 ) 2 ( 0 . 051 ) 1 ( 0 . 083 ) 2 ( 0 . 080 ) 1 ( 0 . 023 ) 5 ( 0 . 161 ) 1 ( 0 . 017 ) fulvescens Niviventer 27 ( 0 . 199 ) 41 ( 1 . 519 ) 35 ( 0 . 294 ) 24 ( 0 . 180 ) 24 194 ( 0 . 051 ) ( 0 . 051 ) Rattus 2 ( 0 . 045 ) 5 ( 0 . 027 ) 8 ( 0 . 067 ) 9 ( 0 . 130 ) 2 ( 0 . 080 ) 8 ( 0 . 094 ) 1 ( 0 . 143 ) remotus 15 63 ( 0 . 051 ) ( 0 . 051 ) Tupaia 4 ( 0 . 022 ) 2 ( 0 . 022 ) 1 ( 0 . 020 ) 1 ( 0 . 026 ) 1 ( 0 . 083 ) 1 ( 0 . 019 ) 1 ( 0 . 014 ) 1 ( 0 . 040 ) 2 ( 0 . 015 ) 7 ( 0 . 082 ) 9 ( 0 . 191 ) 1 ( 0 . 032 ) 1 ( 0 . 017 ) belangeri 15 ( 0 . 146 ) 21 ( 0 . 176 ) 29 121 ( 0 . 051 ) ( 0 . 051 ) 2 ( 0 . 111 ) 1 ( 0 . 010 ) 2 ( 0 . 022 ) 4 ( 0 . 029 ) 4 ( 0 . 148 ) 8 ( 0 . 067 ) 3 ( 0 . 058 ) 2 ( 0 . 029 ) 6 ( 0 . 240 ) 4 ( 0 . 200 ) 8 ( 0 . 060 ) 6 ( 0 . 071 ) 1 ( 0 . 021 ) 2 ( 0 . 143 ) 2 ( 0 . 032 ) 2 ( 0 . 034 ) Menetes 11 ( 0 . 059 ) 34 berdmorei 527 ( 0 . 051 ) ( 0 . 051 ) 7 ( 0 . 038 ) 1 ( 0 . 010 ) 1 ( 0 . 011 ) 1 ( 0 . 007 ) 1 ( 0 . 056 ) 1 ( 0 . 050 ) 1 ( 0 . 016 ) 2 ( 0 . 034 ) finlaysonii 20 ( 0 . 455 ) 11 ( 0 . 092 ) 13 ( 0 . 098 ) 20 Callosciurus 106 7 18 44 12 37 91 50 31 39 27 12 18 52 26 69 25 20 85 47 14 62 43 31 73 59 186 103 136 119 133 Day sp. sp. sp. 3 Speceis Name Michelia baillonii Aglaia lawii Aglaia spectabilis Aphanamixis polystachya Dysoxylum cyrtobotorium Melia azedarach Sandoricum koetjape Antiaris toxicaria Artocarpus lakoocha Ficus altissima Ficus annulata Ficus benjamina Ficus subcoradata Ficus Knema erratica Syzygium albiflorum Syzygium cumini Areca triandra Livistona speciosa Helicia formosana Anthocephalus chinensis Mitragyna Canthium coffeoides Acronychia pedunculata Citrus limon Citrus Clausena harmandiana Mischocarpus pentapetalus Palaquim garetti Aphananthe cuspidata No. of plant species visited No. of visit Family Magnoliaceae Meliaceae Moraceae Myristicaceae Myrtaceae Palmae Proteaceae Rubiaceae Rutaceae Sapindaceae Sapotaceae Ulmaceae . Continued. Table 3 . The number of visits per day is shown in parenthesis. Day: the number of days when photographing was made. 26 Shunsuke SUZUKI, Shumpei KITAMURA, Masahiro KON, Pilai POONSWAD, Phitaya CHUAILUA, Kamol PLONGMAI, Takakazu YUMOTO, Naohiko NOMA, Tamaki MARUHASHI and Prawat WOHANDEE

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Received 27th July 2005 Accepted 24th Jan. 2006 28 Shunsuke SUZUKI, Shumpei KITAMURA, Masahiro KON, Pilai POONSWAD, Phitaya CHUAILUA, Kamol PLONGMAI, Takakazu YUMOTO, Naohiko NOMA, Tamaki MARUHASHI and Prawat WOHANDEE 5 . 6 6 . 8 9 . 3 9 . 3 7 . 4 23 23 23 11 . 1 10 . 3 13 . 2 5 . 6 6 . 5 8 . 4 7 . 4 4 . 8 . 5 22 22 22 15 . 4 11 . 1 5 . 6 9 . 7 3 . 7 9 . 4 21 21 21 10 . 2 10 . 3 11 . 1 11 . 0 5 . 6 7 . 5 . 6 7 . 4 9 . 20 20 20 13 . 0 12 . 4 11 . 3 8 . 3 7 . 9 . 3 9 . 19 19 19 33 . 3 14 . 3 22 . 2 14 . 8 1 . 3 0 . 9 0 . 9 2 . 5 8 . 3 9 . 8 8 . 3 1 . 9 8 . 5 1 . 9 . 4 3 . 2 18 18 18 10 . 3 12 . 7 2 . 6 1 . 2 1 . 4 1 . 5 2 . 8 0 . 1 17 17 17 17 . 6 11 . 6 15 . 7 25 . 2 16 . 7 15 . 9 15 . 5 16 . 0 4 . 2 . 4 9 . 6 9 . 4 6 . 5 8 . 3 3 . 7 5 . 6 5 . 7 16 16 16 10 . 4 22 . 3 12 . 1 13 . 2 10 . 3 5 . 9 3 . 7 6 . 3 5 . 6 4 . 9 5 . 6 4 . 7 15 15 15 14 . 3 10 . 8 10 . 2 17 . 8 13 . 6 13 . 2 15 . 1 1 . 5 5 . 9 2 . 8 4 . 3 2 . 8 1 . 2 1 . 9 14 14 14 14 . 3 13 . 3 19 . 2 18 . 2 17 . 0 17 . 0 4 . 4 . 3 5 . 6 0 . 8 5 . 6 6 . 1 5 . 6 5 . 7 13 13 13 10 . 4 15 . 7 13 . 7 10 . 6 11 . 3 10 . 4 7 . 8 1 . 5 9 . 6 0 . 5 5 . 9 . 1 9 . 4 7 . 5 2 . 8 1 . 8 4 . 2 1 . 2 1 . 4 0 . 9 12 12 12 6 . 5 9 . 6 2 . 2 . 7 5 . 7 1 . 9 1 . 5 4 . 2 1 . 2 1 . 9 11 11 11 Time of day 9 . 1 7 . 4 6 . 0 2 . 6 . 8 7 . 6 6 . 4 . 6 1 . 5 5 . 6 6 . 1 7 . 0 4 . 7 10 10 10 11 . 3 9 9 9 5 . 2 7 . 2 2 . 2 . 7 6 . 1 3 . 8 5 . 7 4 . 0 0 . 0 . 9 8 8 8 9 . 1 1 . 5 4 . 8 4 . 0 8 . 2 8 . 5 2 . 8 5 . 8 4 . 2 1 . 2 1 . 4 0 . 9 10 . 6 11 . 3 7 7 7 7 . 8 8 . 9 . 6 6 . 7 8 . 2 9 . 1 7 . 5 7 . 5 7 . 3 9 . 7 7 . 0 12 . 0 11 . 0 14 . 2 6 6 6 1 . 3 8 . 4 7 . 3 4 . 1 1 . 5 1 . 9 2 . 8 33 . 8 32 . 4 20 . 9 23 . 6 36 . 6 36 . 6 31 . 1 5 5 5 2 . 9 1 . 2 7 . 4 3 . 0 . 9 0 . 5 1 . 4 3 . 7 2 . 4 1 . 4 6 . 1 . 9 3 . 5 19 . 6 4 4 4 1 . 7 . 4 9 . 7 9 . 3 9 . 3 4 . 9 . 0 11 . 1 12 . 8 3 3 3 8 . 3 9 . 5 7 . 4 9 . 0 16 . 7 10 . 3 13 . 0 11 . 0 2 2 2 5 . 6 7 . 4 7 . 3 9 . 3 7 . 7 . 8 10 . 3 13 . 0 1 1 1 5 . 6 9 . 3 7 . 8 7 . 4 8 . 1 10 . 3 13 . 0 13 . 2 0 0 0 5 . 6 5 . 1 8 . 4 7 . 4 9 . 3 7 . 7 . 9 10 . 2 77 68 83 73 18 66 53 39 54 72 54 82 71 91 372 108 106 108 397 106 N N N 1108 1418 visitations by both mammal species for a given pair. C. finlaysonii belangeri T. C. finlaysonii M. berdmorei C. finalysonii R. remotus C. finalysonii N. fulvescens C. finalysonii L. sabanus C. finalysonii M. surifer belangeri T. M. berdmorei belangeri T. R. remotus belangeri T. L. sabanus belangeri T. N. fulvescens belangeri T. M. surifer Appendix. Comparison of Appendix. the proportion (%) of visits for each hour period between each pair of small mammal species. N, the total number of fruit Fruit visitation patterns of small mammals in Thailand 29 9 . 6 8 . 5 7 . 5 0 . 7 . 6 . 8 9 . 3 7 . 6 8 . 6 6 . 7 23 23 23 10 . 4 13 . 5 11 . 1 13 . 3 12 . 5 6 . 3 7 . 2 8 . 9 3 . 8 8 . 9 7 . 8 . 3 . 8 . 2 9 . 0 9 . 0 22 22 22 12 . 8 14 . 3 11 . 5 10 . 0 10 . 0 2 . 1 9 . 6 9 . 2 7 . 1 5 . 8 9 . 5 9 . 8 9 . 5 8 . 6 9 . 7 21 21 21 10 . 6 11 . 5 11 . 5 11 . 1 10 . 0 10 . 0 4 . 2 8 . 5 9 . 6 7 . 20 20 20 12 . 0 12 . 0 11 . 5 13 . 3 11 . 7 11 . 7 10 . 0 11 . 3 11 . 4 12 . 9 12 . 4 6 . 7 7 . 5 7 . 1 19 19 19 22 . 9 11 . 4 10 . 6 13 . 8 28 . 6 11 . 5 23 . 1 21 . 2 14 . 7 11 . 7 13 . 4 12 . 5 12 . 9 9 . 7 2 . 1 1 . 8 9 . 0 6 . 9 3 . 1 1 . 9 3 . 4 1 . 7 1 . 5 3 . 1 3 . 2 18 18 18 13 . 9 0 . 1 0 . 1 17 17 17 21 . 6 27 . 1 24 . 6 24 . 6 9 . 5 9 . 0 16 16 16 11 . 9 11 . 1 7 . 3 5 . 6 6 . 4 . 8 15 15 15 4 . 6 3 . 7 4 . 3 3 . 8 14 14 14 0 . 3 0 . 7 0 . 9 0 . 8 13 13 13 1 . 8 1 . 7 2 . 0 1 . 7 12 12 12 1 . 8 1 . 5 1 . 4 1 . 5 11 11 11 Time of day 1 . 8 2 . 0 1 . 7 2 . 3 10 10 10 9 9 9 4 . 6 3 . 4 4 . 3 3 . 4 8 8 8 6 . 4 6 . 1 6 . 9 6 . 3 7 7 7 8 . 5 6 . 8 8 . 0 6 . 1 6 6 6 0 . 1 0 . 1 0 . 1 19 . 5 22 . 2 20 . 9 20 . 8 5 5 5 0 . 3 4 . 2 7 . 2 0 . 7 0 . 6 1 . 0 3 . 7 7 . 1 5 . 8 3 . 8 3 . 8 3 . 6 5 . 0 7 . 2 3 . 7 3 . 8 4 4 4 8 . 3 9 . 6 9 . 2 7 . 1 5 . 8 7 . 8 . 9 7 . 9 . 1 5 . 0 8 . 9 . 7 10 . 6 12 . 5 10 . 0 10 . 0 3 3 3 7 . 2 8 . 5 8 . 7 7 . 1 7 . 6 . 7 9 . 6 9 . 4 7 . 5 7 . 2 8 . 6 12 . 5 11 . 5 13 . 3 10 . 1 10 . 4 2 2 2 7 . 2 7 . 7 . 1 7 . 7 . 2 7 . 4 7 . 1 10 . 4 12 . 8 11 . 5 11 . 5 15 . 6 11 . 5 10 . 0 12 . 5 14 . 3 1 1 1 8 . 3 9 . 6 8 . 3 7 . 1 9 . 6 3 . 8 7 . 7 . 1 9 . 3 8 . 4 7 . 2 10 . 6 11 . 1 11 . 7 12 . 5 12 . 9 0 0 0 8 . 3 7 . 2 6 . 4 7 . 3 . 8 7 . 6 . 7 9 . 6 8 . 1 3 . 5 . 0 6 . 7 7 . 4 7 . 1 7 . 14 . 3 48 47 14 52 26 45 52 60 40 70 329 410 166 350 524 N N N 928 194 834 1577 1065 M. berdmore R. remotus M. berdmore N. fulvescen M. berdmore L. sabanu M. berdmore M. surife R. remotus N. fulvescen R. remotus L. sabanu R. remotus M. surife N. fulvescen L. sabanu N. fulvescen M. surife L. sabanu M. surife Appendix. Continued Appendix.